Effects of leaf and sap feeding insects on photosynthetic rates of goldenrod

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Abstract
Herbivory can alter the balance between sources and sinks within a plant, and changes in the source-sink ratio often lead to changes in plant photosynthetic rates. We investigated how feeding by three insect herbivores affected photosynthetic rates and growth of goldenrod (Solidago altissima). One, a phloem-sap feeding aphid (Uroleucon caligatum), creates an additional sink, and the other two, a leaf-chewing beetle (Trirhabda sp.) and a xylem-sap feeding spittlebug (Philaenus spumarius) both reduce source supply by decreasing leaf area. Plants were grown outside in large pots and insects were placed on them at predetermined densities, with undamaged plants included as controls. All insects were removed after a 12-day feeding period. We measured photosynthetic rates both of damaged leaves and of undamaged leaves that were produced after insect removal. Photosynthetic rates per unit area of damaged leaves were reduced by spittlebug feeding, but not by beetle or aphid feeding. Conductance of spittlebugdamaged leaves did not differ from controls, but internal carbon dioxide concentrations were increased. These results indicate that spittlebug feeding does not cause stomatal closure, but impairs fixation within the leaf. Effects of spittlebug feeding on photosynthetic rates persisted after the insects were removed from the plants. Photosynthetic rates per unit area of leaves produced after insect removal on spittlegug-damaged plants were lower than control levels, even though the measurements were taken 12 days after insect removal. The measurement leaf on spittlebugdamaged plants was reduced in area by 27% relative to the controls, but specific leaf area (leaf area/leaf weight) was increased by 18%. Because of the shift in specific leaf area, photosynthetic rates were also examined per unit leaf weight, and when this was done there were no significant differences between control and spittlebug-damaged plants. Beetle and aphid feeding had no effects on the photosynthetic rate of the leaves produced after insect removal. Plant relative growth rates (in terms of height) were reduced by spittlebugs during the period that the insects were feeding on the plants. Relative growth rates of spittlebug-damaged plants were increased above control levels after insect removal, but these plants were still shorter than controls 17 days after insect removal. Beetles and aphids did not affect plant relative growth rates or plant height. Feeding by both spittlebugs and beetles reduced leaf area, and the effect of the spittlebug was more severe than that of the beetle. These results show that effects of herbivory on photosynthetic rates cannot be predicted simply by considering changes in the source-sink ratio, and that spittlebug feeding is more damaging to the host plant than beetle or aphid feeding.